Gerald D. Schmidt

Acanthocephalan infections of man, with two new records.

Previous records of acanthocephalans in man are reviewed. Acanthocephalus bufonis (Shipley, 1903) Southwell et Macfie, 1925, is reported from man for the first time, recovered on autopsy of an Indonesian in Djakarta. Corynosoma strumosum (Rudolphi, 1802) Liihe, 1904, is reported from an Eskimo in Alaska, the first record of this parasite from man. Infections of man with acanthocephalan parasites are rare. Many species in this phylum are known to have low host specificity (Golvan, 1957) but for ecological reasons are not likely to be ingested by humans. The intermediate hosts of all species whose life cycles are known are either crustaceans, myriopods, or insects. However, several species are known to utilize fish, amphibians, or reptiles as paratenic hosts, thereby increasing the chances of accidental parasitism in persons eating such animals raw or undercooked. Lambl (1859) gave the first report of a thorny-headed worm in man when he recovered a Macracanthorhynchus hirudinaceus from a child in Prague. Lindemann (1865) stated the infection was common in Russia, where Schneider (1871) found that scarabaeid beetle grubs, the intermediate host, were commonly eaten raw. Gonzaga (1921) recovered eggs of M. hirudinaceus in two of 1,236 human stools in Brazil. A more recent case of proved chronicity was published by Pradatsundarasar and Pechranond (1965) in Thailand. Moniliformis moniliformis, a parasite of rats and cockroaches, has been reported from man in Italy, Sudan, and British Honduras (Faust and Russell, 1957), and Beck (1959) reported it from a patient in Florida. Grassi and Calandruccio (1888) reported on the symptoms displayed by the second author after voluntarily infecting himself with M. moniliformis. Kuntz (1960) reported acanthocephalan eggs, possibly those of this species, from stools of two children in Dacca, East Pakistan. Moore, Fry, and Englert (1969) made the interesting discovery of acanthocephalan eggs in human coprolites obtained in archeological excavations in Danger Cave, Utah. RadiocarReceived for publication 20 October 1970. bon dating placed the samples at 1869 BC ? 160 years, and AD 20 ? 240 years. The authors suggested that the parasite may have been Moniliformis clarki, which is common in the area. The only acanthocephalan outside the class Archiacanthocephala reported from man is A anthocephalus rauschi, recovered by Dr. Rob rt Rausch from the peritoneum of an Eskimo in Alaska, and described by Golvan (1969). The normal definitive host is probably a fish. The present paper reports the occurrence of two more palaeacanthocephalans in humans. Acanthocephalus bufonis (Shipley, 1903) Southwell et Macfie, 1925 (Syn. Acanthocephalus sinensis Van Cleave, 1937) In 1954, during a routine autopsy by Dr. Lie Kian Joe at the Medical School in Djakarta, Indonesia, four male acanthocephalans were recovered from the small intestine of an Indonesian man by screening the contents of the intestine with a metal sieve routinely used to collect hookworms. No wild animals were dissected in the autopsy room, and there was no chance that the specimens came from an outside source and had become mixed with material from human autopsies. Due to a lack of library facilities in Djakarta it was impossible to identify the worms to species; they were reported as Acanthocephalus sp. by Lie Kian Joe and Tan Kok Siang (1959). One specimen was lost prior to the present study. I now know them to be adult males of Acanthocephalus bufonis, a very common parasite of amphibians in Asia (Yuen and Fernando, 1967). Its life cycle is unknown but probably involves a terrestrial isopod crustacean as intermediate host. I can speculate that the man became infected in one of three ways: (1) by ingesting an infected arthropod; (2) by eating an am-

Nematode parasites of Oceanica: IX. Capillariids from Passeriform hosts collected in Taiwan

The capillariid nematodes present in passeriform hosts collected in Taiwan are recorded and described. Two new species are established, Capillaria parusi sp.n. from Parus monticolus insperatus and Sitta europaea formosana , and C. madseni sp.n. from Corvus macrorhynchus colonorum , and their affinities are discussed. Capillaria tridens, C. longifila and C. contorta are recorded from a number of new hosts.

Resurrection of Southwellina Witenberg, 1932, with a description of Southwellina Dimorpha sp. n., and a key to genera in Polymorphidae (Acanthocephala).

The family Polymorphidae is briefly discussed, and a key to its genera is presented. The subfamilies Polymorphinae Meyer, 1931, and Corynosomatinae Petrochenko, 1956, are suppressed. Southwellina Witenberg, 1932, is considered valid and is redefined. It contains S. hispida (Van Cleave, 1925), S. macracanthus (Ward et Winter, 1952) comb. n., and S. dimorpha sp. n. The new species was found as juveniles in crayfish, Procambarus clarkii, in Louisiana, and as adults in white ibis, Eudocimus albus, in Florida. It differs from other species in its sexual dimorphism, the female having large hooks and 1 field of trunk spines, and the male having smaller hooks and 2 fields of trunk spines. The following species are considered synonyms of S. hispida: Arhythmorhynchus duocinctus Chandler, 1935; A. fuscus Harada, 1939; Polymorphus ardeae Belopolskaia, 1958; and Hemiechinosoma ponticum Petrochenko et Smogorjevskaia, 1962. Skrjabinorhynchus Petrochenko, 1956, is suppressed and its species are recombined as Arhythmorhynchus capellae (Yamaguti, 1935) comb. n., and Arhythmorhynchus eroliae (Yamaguti, 1939) comb. n. The resulting homonym, A. capellae Schmidt, 1963, is renamed Arhythmorhynchus jeffreyi nom. n. The family Polymorphidae Meyer, 1931, is a homogeneous assemblage of species, most of which are parasites of aquatic birds and mammals. Distinction between species is usually difficult, and between genera even more so. When first proposed, most genera were based on one or a few species. Each genus had some distinctive characteristics which readily separated it from other genera. But as more species were discovered, differences between genera became less apparent; today very slight distinctions are used to separate them. For example, Corynosoma differs from Polymorphus only in possessing genital spines, and in some species the spines are on only one sex. Efforts by Petrochenko (1956) and Yamaguti (1963) to divide the family into the subfamilies Polymorphinae Meyer, 1931, and Corynosomatinae Petrochenko, 1956, resulted in placing genera as obviously closely related as Polymorphus and Corynosoma in different subfamilies. As this is less than useful, I cannot accept these subfamilies as valid. The discovery of a new species in this family has stimulated me to review the status of these related genera. It could be logically argued that all species in this family belong to a single genus. On the basis of similarity of form, embryology, and biology they are obviously genetically similar. However, for a Received for publication 13 October 1972. classification to be workable it must help distinguish between species groups in the most convenient manner possible. Therefore, I have taken a more liberal approach and recognize nin genera, which can be diagnosed by the following key. Key to genera in Polymorphidae Meyer, 1931 (Syn. Filicollidae Petrochenko, 1956) la. Cement glands kidney-shaped. Proboscis of adult female greatly swollen, with hooks restricted to short, radially arranged rows on anterior surface. Parasites of water birds Filicollis Liihe, 1911 lb. Cement glands claviform or tubular. Proboscis swollen or not; when swollen, longitudinal rows of hooks cover entire proboscis 2 2a. Genital spines present on at least one sex. Parasites of aquatic birds and mammals --.---... ---Corynosoma Liihe, 1904 (Syn. Chentrosoma Monticelli, 1905, nec Porta, 1906; Echinosoma Porta, 1907, partim.) 2b. Gental spines absent on both sexes ----3 3a. Anterior end of trunk greatly swollen, separated from rest of trunk by conspicuous constriction 4 3b. Anterior trunk not separated from posterior portion by narrow constriction 5 4a. Anterior trunk with 1 field of spines surrounding it. Parasites of Cetacea .....--Bolbosoma Porta, 1908 (Syn. Echinorhynchus Zoega in Miiller, 1776, partim; Bolborhynchus Porta, 1906.)

Revision of the class Archiacanthocephala Meyer, 1931 (phylum Acanthocephala), with emphasis on Oligacanthorhynchidae Southwell et Macfie, 1925

Class Archiacanthocephala is modified to include Moniliformida ord. n. All genera in Oligacanthorhynchidae were studied and several changes in classification of the family are made. Nephridiacanthus Meyer, 1931; Hamanniella Travassos, 1915; Travassosia Meyer, 1931; and Echinopardalis Travassos, 1918, are considered to be junior synonyms of Oligacanthorhynchus Travassos, 1915. Prosthenorchis elegans (Diesing, 1851) Travassos, 1915, P. lemuri Machado, 1950, and P. fraterna (Baer, 1959) comb. n. are retained in Prosthenorchis Travassos, 1915: all other species previously assigned to this genus are distributed among Pachysentis Meyer, 1931, Oncicola Travassos, 1916, and Neoncicola gen. n. The new genus is proposed for 7 species similar to Oncicola but with 30 rather than 36 proboscis hooks. Keys are provided for the orders in class Archiacanthocephala, and the 8 genera in Oligacanthorhynchidae accepted as valid. Each genus is redefined and a list of species in each is provided. No family in the Acanthocephala has a more confused literature than Oligacanthorhynchidae Southwell et Macfie, 1925. This condition has resulted from a number of causes: vague descriptions of species and uncertain definitions of genera by early authors, incorrect interpretations of morphology by more recent authors, and the homogeneity of the species themselves. Consequently it is almost impossible to identify an oligacanthorhynchid to genus, even using the keys of Petrochenko (1958), Golvan (1962), and Yamaguti (1963). During the past 10 years I have accumulated a large series of specimens representing all of the genera in the family. Several collectors have contributed specimens for study, most notably Dr. John F. Schacher who generously gave me a large collection which he assembled in Lebanon from 1961 to 1963. Study of these collections has made it possible to revise the family on sound morphological lines, without resorting to the separation of genera on the basis of definitive hosts, as has been historically accepted (e.g., Oligacanthorhynchus from birds, Hamanniella from opossums and edentates, etc.). Despite my efforts, and the efforts of others, it remains impossible to identify most juvenile forms to genus. The monotonous similarity of proboscis armature between most genera and species forces the systematist to rely on adult Received for publication 15 July 1971. characters, such as location of testes and shape of trunk, features that are unresolved in juveniles. The practice of describing juveniles as new species within a given genus in this family should be avoided, for it only continues to confound an already confused classification. The lists of species following the diagnoses of g nera in this paper have, for the most part, been uncritically drawn from the literature with little consideration of possible synonyms within each list. However, each species poorly known or known from juveniles only appears with a question mark before it. My interest here has been to resolve problems at the levels of genus and above; this should simplify the task for others who wish to further consider the species in each genus. In addition to studying whole mounts, stained with Semichons carmine, I employed microdissections and serial sectioning to resolve certain problems, such as structure and insertions of the muscles of the proboscis receptacles. The typically thick walls of the trunk and the sheer size of many species make studies of whole mounts less practical than with most other families of Acanthocephala. The practice of describing proboscis armature as forming spiral rows of hooks has been avoided in this paper. The difficulties of counting hooks by this technique outweigh any advantage of ease of description. Instead, I consider the hooks to be arranged in short, approximately longitudinal rows.

Life Cycle and Development of Prosthorhynchus formosus (Van Cleave, 1918) Travassos, 1926, an Acanthocephalan Parasite of Birds

The life cycle of Prostlwrhynchus formosus (Van Cleave, 1918) Travassos, 1926 is presented. The terrestrial isopods Amwdillidium vulgare, Porcellio laevis, and P. scaber served as experimental intermediate hosts, and chickens and turkeys as experimental definitive hosts. After ingestion of the eggs by the isopod, the acanthor emerges from its shells within 15 min to 2 hr, enters the gut wall of its host, and remains there 15 to 25 days. It then migrates to the hemocoel and develops through the acanthella stage to the infective cystacanth by the 60th to 65th day of infection . Van Cleave (1918) described Plagiorhynchus formosus on the basis of four specimens collected from the flicker, Colaptes auratus, at Bowie, Maryland. In 1926, Travassos reassigned this species to the genus Prosthorhynchus but gave no reasons for the change. Van Cleave (1942) disagreed strongly with this change in generic status and reaffirmed its position in Plagiorhynchus. Golvan (1956) clarified the differences between Plagiorhynchus and Prosthorhynchus and defended its position in Prosthorhynchus. This view, which was upheld by Petrochenko (1956) and Yamaguti (1963), is accepted by the present authors. Jones (1928) extended the host list to include the chicken (Gallus domesticus) , robin (Turdus migratorius) , and the crow (Corvus americanus). She remarked on the possibilities of this worm becoming an important parasite of domestic fowl. Cuvillier (1934) added the catbird (Dumatella carolinensis), and a thrush Received for publication 17 April 1964. ... Part of a thesis submitted by the senior author to the Graduate School of Colorado State University in partial fulfillment of the requirements for the Ph.D. degree, February 1964. This investigation was supported by Training Grant PHS-2E-94 (C2,3) from the National Institute of Allergy and Infectious Diseases of the National Institutes of Health, U. S. Public Health Service, and by a fellowship from the Boettcher Foundation, Denver, Colorado. 721 (Hylocichla sp.), and Van Cleave (1942) the towhee (Pipilo erythrophthalmus) , starling (Sturnus vulgaris), grackle (Quiscalus quiscala) , and hermit thrush (H ylocichla guttata). Chandler and Rausch (1949) found it in the brown thrasher (T oxostoma rufum) , and Hunter and Quay (1953) in Macgillivrays seaside sparrow (Ammospiza maritima macgillivaraii). The present report adds the red-shafted flicker (Colaptes cafer) and the domestic turkey (experimental) . Sinitsin (1929) reported finding a juvenile of Plagiorhynchus formosus in the sow bug, Armadillidium vulgare, near Washington, D. C. To date, this is the only report on the life history of P. formosus, although Dollfus and Dalens (1960) reported larvae of Prosthorhynchus cylindraceus in A. vulgare in France. This work presents the complete life cycle of Prosthorhynchus formosus as observed under laboratory conditions, utilizing three species of terrestrial isopods as intermediate hosts and domestic fowl as definitive hosts . MATERIALS AND METHODSThe life cycle of Prosthorhynchus formosus (Van Cleave, 1918) Travassos, 1926 is presented. The terrestrial isopods Armadillidium vulgare, Porcellio laevis, and P. scaber served as experimental intermediate hosts, and chickens and turkeys as experimental definitive hosts. After ingestion of the eggs by the isopod, the acanthor emerges from its shells within 15 min to 2 hr, enters the gut wall of its host, and remains there 15 to 25 days. It then migrates to the hemocoel and develops through the acanthella stage to the infective cystacanth by the 60th to 65th day of infection. Van Cleave (1918) described Plagiorhynchus formosus on the basis of four specimens collected from the flicker, Colaptes auratus, at Bowie, Maryland. In 1926, Travassos reassigned this species to the genus Prosthorhynchus but gave no reasons for the change. Van Cleave (1942) disagreed strongly with this change in generic status and reaffirmed its position in Plagiorhynchus. Golvan (1956) clarified the differences between Plagiorhynchus and Prosthorhynchus and defended its position in Prosthorhynchus. This view, which was upheld by Petrochenko (1956) and Yamaguti (1963), is accepted by the present authors. Jones (1928) extended the host list to include the chicken (Gallus domesticus), robin (Turdus migratorius), and the crow (Corvus americanus). She remarked on the possibilities of this worm becoming an important parasite of domestic fowl. Cuvillier (1934) added the catbird (Dumatella carolinensis), and a thrush Received for publication 17 April 1964. * Part of a thesis submitted by the senior author to the Graduate School of Colorado State University in partial fulfillment of the requirements for the Ph.D. degree, February 1964. This investigation was supported by Training Grant PHS-2E-94 (C2,3) from the National Institute of Allergy and Infectious Diseases of the National Institutes of Health, U. S. Public Health Service, and by a fellowship from the Boettcher Foundation, Denver, Colorado. (Hylocichla sp.), and Van Cleave (1942) the towhee (Pipilo erythrophthalmus), starling (Sturnus vulgaris), grackle (Quiscalus quiscala), and hermit thrush (Hylocichla guttata). Chandler and Rausch (1949) found it in the brown thrasher (Toxostoma rufum), and Hunter and Quay (1953) in Macgillivrays seaside sparrow (Ammospiza maritima macgillivaraii). The present report adds the red-shafted flicker (Colaptes cafer) and the domestic turkey (experimental). Sinitsin (1929) reported finding a juvenile of Plagiorhynchus formosus in the sowbug, Armadillidium vulgare, near Washington, D. C. To date, this is the only report on the life history of P. formosus, although Dollfus and Dalens (1960) reported larvae of Prosthorhynchus cylindraceus in A. vulgare in France. This work presents the complete life cycle of Prosthorhynchus formosus as observed under laboratory conditions, utilizing three species of terrestrial isopods as intermediate hosts and domestic fowl as definitive hosts. MATERIALS AND METHODS

Revision of Mediorhynchus van cleave 1916 (acanthocephala) with a key to species.

Species characters in Mediorhynchus are discussed, and a consistent method of describing hook arrangement, longitudinal rows, is proposed. Mediorhynchus papillosus Van Cleave 1916, and M. orientalis Belopolskaya 1953 are redescribed, and M. edmondsi sp. n. is described from Palawan. The new species is most similar to M. rodensis Cosin 1971 in armature, but differs from it in proboscis shape and overall size. Among other new records are M. gallinarum (Bhalerao, 1937) in chickens in Palawan, and M. turnixena (Tubangui, 1933) from a button quail in Palawan. Several synonymies are pointed out and several unidentifiable species are mentioned. A key to 29 species recognized as valid is provided.

Nematode parasites of Puerto Rican tree frogs, Eleutherodactylus spp: two new species and a proposal of Poekilostrongylus gen. nov. (Trichostrongylidae).

Poekilostrongylus puertoricensis gen.nov., sp.nov. is proposed for nematodes recovered from Eleutherodactylus coqui, in Puerto Rico. The new genus is similar to Oswaldocruzia Travassos, 1917, but lacks longitudinal ridges on the cuticle. Oswaldocruzia lenteixierai Viqueras, 1938, is partially redescribed, and a key to the genera of the Oswaldocruziinae is given. Thelandros (Parathelandros) garciai sp.nov. is described from E. antillensis and E. portoricensis. Strongyloides sp. was found in E. antillensis and Aplectana spp. (females) were recovered from E. locustus, E. richmondi and E. coqui.

Andracantha, a new genus of Acanthocephala (Polymorphidae) from fish-eating birds, with descriptions of three species.

Problems in taxonomy of the Polymorphidae are discussed, with particular reference to trunk spines. Andracantha gen. n. is proposed for species with genital spines and 2 fields of trunk spines. Corynosoma gravida Alegret 1941, C. mergi Lundstroöm 1941 and C. phalacrocoracis Yamaguti 1939 are redescribed and placed in Andracantha, with A. gravida (Alegret, 1941) comb. n. designated as type species.

SCLEROCOLLUM RUBRIMARIS GEN. ET SP. N. (RHADINORHYNCHIDAE: GORGORHYNCHINAE), AND OTHER ACANTHOCEPHALA OF MARINE FISHES FROM ISRAEL

Sclerocollum rubrimaris gen. et sp. n. from Siganus spp. differs from others in Rhadinorhynch- idae in lacking trunk spines but possessing irregular plaques of hook material in the wall of the anterior trunk. Sclerocollum robustus (Edmonds 1964) comb. n. is included in the new genus. Arhythmacanthus overstreeti sp. n., found in Silhouettea insinuans, differs from all other species in the genus in numbers and sizes of proboscis hooks and in having spines over the entire trunk. Arhythmacanthus plotosi (Yam. 1955) comb. n. is removed from Heterosentis. Golvanacanthus blennii Paggi et Orecchia 1972 is reported from several new host species, and its hooks are redescribed. Other species recorded are Neoechino- rhynchus agilis (Rud.), N. coiliae Yam., and Filosoma sp.

Tapeworms of the Chaco Boreal, Paraguay, with two new species.

Mathevotaenia paraguayae sp. n. is described from Euphractus sexcinctus. It differs from M. didelphidis (Rud.) in having a transverse cirrus pouch anterior to the vagina. Monoecocestus mackiewiczi sp. n. was found in Phyllotis sp. It differs from M. parcitesticulatus Rego in having a much smaller scolex, a poral ovary, and a cirrus pouch 360 to 440 micrometers long. Other species reported are Taenia macrocystis, Taenia omissa, Spirometra mansonoides, Spirometra gracilis, Spirometra longicollis, Diphyllobothrium trinitatis, Atriotaenia parva, and Mathevotaenia tetragonocephala.